A New South Brazilian Species of Actinostemon (Euphorbiaceae)

Transcrição

A New South Brazilian Species of Actinostemon (Euphorbiaceae)
A New South Brazilian Species of Actinostemon (Euphorbiaceae)
Author(s): Luciana dos Santos Dias de Oliveira, André Laurênio de Melo, Marcos José da Silva, Paula
Pinto Eymael, and Margareth Ferreira de Sales
Source: Systematic Botany, 40(2):522-526.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364415X688709
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Systematic Botany (2015), 40(2): pp. 522–526
© Copyright 2015 by the American Society of Plant Taxonomists
DOI 10.1600/036364415X688709
Date of publication August 10, 2015
A New South Brazilian Species of Actinostemon (Euphorbiaceae)
Luciana dos Santos Dias de Oliveira,1,4 André Laurênio de Melo,2 Marcos José da Silva,3
Paula Pinto Eymael,1 and Margareth Ferreira de Sales1
1
Programa de Pós-Graduação em Botânica, Departamento de Biologia, Universidade Federal Rural de Pernambuco,
52171–900, Recife, PE, Brazil.
2
Unidade Acadêmica de Serra Talhada, Universidade Federal Rural de Pernambuco, 56900–000, Serra Talhada, PE, Brazil.
3
Instituto de Ciências Biológicas, Departamento de Botânica, Universidade Federal de Goiás, 74001–970, Goiânia, GO, Brazil.
4
Author for correspondence ([email protected])
Communicating Editor: Min Feng
Abstract—A new species of Actinostemon is described and illustrated. Actinostemon roselii is mentioned only for the states of Paraná
and Santa Catarina (Brazil), where it grows in riparian tropical rainforests. This species can be recognized by its conspicuous and depressed
ovate bracts of the staminate cymule, foliar buds globoid, staminate cymule with three or four flowers, and staminate and pistillate flowers
monochlamydeous. Comments about its distribution and ecology are provided, as well as a key to identify the Brazilian species of Actinostemon.
Based on IUCN criteria, A. roselii can be considered a data deficient (DD) species.
Keywords—Actinostemon roselii, Atlantic domain, Hippomaneae, IUCN, taxonomy.
Actinostemon Mart. ex Klotzsch, together with Gymnanthes
Sw., has the most complex taxonomy of the Hippomaneae
tribe due to poorly understood intra- and intergeneric limits.
Additionally, the recognition of the species is hindered by
several nomenclatural and typing problems and by homogeneity of morphological characters, especially floral features
(Esser 2001, 2012; Oliveira 2014). The genus encompasses
approximately 15 Neotropical species and its center of diversity is in Brazil (Esser 2012).
It is characterized by monoecious species; shrubs to small
trees; leaf margins always entire and glands scattered on the
abaxial surface; inflorescences racemose-cymose, thyrsoid or
rarely paniculiform, bisexual or unisexual, preceded by conspicuous basal cataphylls which are imbricate and generally
striate; staminate cymules subtended by a bract which is
biglandular or not; staminate and pistillate sepals reduced
or absent; and carunculate seeds (Esser 2012; Eymael 2012).
As part of the ongoing review of Gymnanthes, the first
author of this work has examined 20 collections of the new
species herein described from Brazilian (ALCB, CGMS, CRI,
FLOR, FUEL, FURB, HBR, ICN, MBM, and RB) and international herbaria (K, M, and Z) (acronyms according to Thiers
[continuously updated]). Most specimens were identified as
Actinostemon concolor (Spreng.) Müll. Arg., the most morphologically similar species. In this study, we discuss geographical
distribution, morphological relationships, conservation status,
and provide an identification key to Actinostemon species occurring in Brazil based on the monograph by Eymael (2012).
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Shrubs to small trees 2–8 m tall, monoecious, glabrous.
Branches cylindrical, lenticels ellipsoid, yellow to blackish.
Foliar buds 1–2 1–2 mm, globoid; cataphylls covering the
buds, more than 1 pair (1.3–2 1–1.5 mm), obovate, imbricate, rigid, with brittle margins, subpersistent. Stipules caducous. Petiole 0.4–1.2 cm long, slender, cylindrical, canaliculate
in the upper part, dark brown to blackish in the voucher
specimens examined. Leaves alternate, regularly distributed
along the branches; leaf blade 3.5–8.5 1.8–4.5 cm, chartaceous
to slightly coriaceous, discolorous, elliptic, elliptic-spatulate,
sometimes obovate, base attenuate, apex acute, sometimes
rounded, margin entire, revolute; glands globoid to ellipsoid,
yellowish to blackish, in variable number in the blade matrix,
sometimes near the base; venation strongly brochidodromous,
secondary veins subpatent, slightly turned to the apex. Inflorescence 1.0–2.5 cm long, thyrsoid, axillary, congested, unisexual staminate or bisexual with 1–2 pistillate flowers proximal,
staminate cymules with 3–4(–5) flowers distal; peduncule short,
ca. 1–2 mm long, with cataphylls (1–1.5 0.8–1.2 mm) slightly
obovate to rounded, imbricate, with no evident veins. Staminate cymule and pistillate flower subtended by a bract.
Staminate bracts 2–2.5 3–3.5 mm, conspicuous, sessile,
membranaceous, yellow-brownish, depressed ovate, slightly
cucullate, apex rounded, rarely acute, reddish, base truncate,
striate, margin entire, sometimes lacerate; gland 1 pair per
bract, basilateral, 0.5–1.2 mm long, ear-shaped, creased, sessile. Staminate flowers distinctly pedicellate, 1.2–4.3 mm
long, generally of unequal size; sepals 1–3, < 0.5 mm long,
disposed along the pedicel at different points of insertion,
membranaceous, hyaline, sometimes achlamydeous; stamens
(2–)3–4(–5), filaments 1–2 mm long, completely free; anthers
dorsifixed, extrorse, rimose. Pistillate bracts 1–2, ca. 1.2–2
1.5–2 mm, deltoid to triangular, membranaceous, margin
entire to slightly erose, sometimes sinuate, hyaline; gland
1 pair per bract, basilateral, sessile, sometimes absent. Pistillate flowers, when in bisexual cymules, flanked by 2 staminate
flowers, or sometimes solitary at the base of the staminate
inflorescence; pedicel 3–3.2 mm long; sepals 3, ca. 1 mm long,
united, valvate, without glands, membranaceous, triangular,
apex acute, margin entire; ovary 1 1 mm, ovoid; styles 1.5–
2 mm long, curved, style column 0.5–1.5 mm long, stigma
surface rugulose. Capsule 5–6 5–8 mm, globoid, smooth,
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Taxonomic Treatment
Actinostemon roselii L. Oliveira, A. L. Melo & M. F. Sales,
sp. nov.—TYPE: BRAZIL. Paraná: Tibagi, Canyon Guartelá,
12 Sep 1996 (fl), C. Giraldi s. n. (holotype: FUEL 21203!;
isotype: K 001096643!).
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Close to Actinostemon concolor (Spreng.) Müll. Arg. due to
leaves distributed along the branches, blade elliptic, and fruits
without appendices, however, it is markedly characterized
by the size of the bracts which precede the staminate cymules
(2–2.5
3–3.5 mm), foliar buds globoid, and presence of
sepals in staminate and pistillate flowers.
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2015]
OLIVEIRA ET AL.: A NEW SOUTH BRAZILIAN SPECIES OF ACTINOSTEMON
523
Fig. 1. Actinostemon roselii L. Oliveira, A. L. Melo & M. F. Sales. A. Habit. B. Flowering branch. C. Detail of foliar bud. D. Foliar glands on the lower
surface. E. Detail of the glands. F. Bisexual inflorescence. G. Staminate cymule. H. Staminate bracts. I. Staminate flowers. J. Bisexual cymule evidencing
the pistillate flower. K. Fruit. L. Columella. M. Detail of the columella with mericarp. N. Detail of the mericarp showing the seeds. O. Carunculate seed.
SYSTEMATIC BOTANY
Additional Specimens Examined—BRAZIL. Paraná: Tibagi, Cânion
Guartelá, 24 300 52.6600 S, 50 240 40.1100 W, 13 Dec 1996 (fr), Silva et al. 1817
(ALCB, CGMS, MBM); same loc., 12 Sep 1996 (fl), Giraldi s. n. (FUEL
21203, K). Santa Catariana: Angelina, Rancho das Tábuas, entorno do
fragmento, 27 370 4900 S, 49 20 5800 W, 590 m, 1 Feb 2010 (fr), Stival-Santos et al.
1629 (FURB, RB); Imaruı́, Serraria Alcides Alves, Águas Mornas,
28 200 42.1000 S, 48 480 58.8700 W, 50 m, 16 Jan 1973 (fr), Klein & Bresolin
10736 (FLOR); Alto Rio D’Uma, 50 m, 21 Sep 1973 (fl), Bresolin 856
(FLOR, ICN); Jacinto Machado, Sanga da Areia, 28 590 56.3800 S,
49 450 48.0000 W, 5 m, 31 Oct 1959 (fr), Reitz & Klein 9318 (HBR, M, Z);
Lauro Müller, Vargem Grande, 28 230 46.1100 S, 49 230 45.1100 W, 400 m,
17 Dec 1958 (fr), Reitz & Klein 8091 (FLOR); Orleans, margin do Rio
Novo, 28 210 25.6700 S, 49 200 6.7400 W, 10 Jan 1992, Martinello & Aguiar
s. n. (CRI 1718); same loc., 28 Nov 1991 (fr), Zanette & Aguiar 1698
(CRI); same loc., 24 Oct 1991 (fl, fr), Zanette et al. 1697 (CRI); same
loc., próxima a Granja Mazon, rio Novo, 14 Nov 1990 (fr), Zanette & Aguiar
991 (CRI); same loc., 12 Apr 1991 (fl), Zanette et al. 1197 (CRI); same
loc., 13 Mar 1992, Zanette et al. 1699 (FLOR); same loc., 12 Sep 1991,
Falkenberg & Zanete 5546 (FLOR); Santo Amaro da Imperatriz, trilha
de turismo ecológico do Hotel Caldas da Imperatriz, 27 410 18.0600 S,
48 460 38.4400 W, 200 m, 28 Nov 1989 (fr), Falkenberg & Alburque 4868
(FLOR); Siderópolis, sı́tio Sete Lombas, 28 350 59.0400 S, 49 250 37.6900 W,
112 m, 12 Sep 2007 (fl), Rodrigues s. n. (CRI 7921).
Taxonomic Notes—Actinostemon was established by
Klotzsch (1841) based on Brazilian material the author named
Actinostemon grandifolius [= A. klotzschii (Didr.) Pax]. Didrichsen
(1857), Grisebach (1857), and Klotzsch (1852) expanded the
circumscription of the genus by describing new species. Müller
(1863, 1866, 1873) proposed several new binomials, trinomials,
and even polynomials for the concept of Actinostemon. Pax and
Hoffmann (1912) and Jablonski (1967) significantly restricted
the circumscription proposed by Müller (1866) accepting
29 and 26 species, respectively. Posteriorly, in his awkward
revision, Jablonski (1969) reduced to 13 the number of
Actinostemon species. Nevertheless, Esser (2012) affirmed that
the revisions performed by Pax and Hoffmann (1912) and
Jablonski (1969) do not reflect the real circumscription of
Actinostemon, since important type collections were not studied and few specimens were analyzed. According to Esser
(2012), Actinostemon comprises about 15 species, whereas in
the World Checklist of Euphorbiaceae (Govaerts et al. 2014)
the genus presents 19 species. Regarding the Brazilian flora,
Oliveira et al. (2013) synonymized Actinostemon unciformis
Jabl. with Gymnanthes klotzschiana Müll. Arg., reducing
the circumscription of the genus. In the Brazilian flora list
(Cordeiro et al. 2014), 16 species of Actinostemon are mentioned. However, the revision of this genus for the country
(Eymael 2012) indicates that Actinostemon presents only
seven species.
The boundaries of Actinostemon species have only been
examined in local floras or regional treatments. The discrepancies in the circumscription of Actinostemon point to
the need for a new monographic revision of the entire
genus, since the number of species recognized by several
authors has varied.
Actinostemon roselii is the first species described in this
genus after the study carried out by Jablonski (1969). This species is clearly differentiated from other species of Actinostemon
mainly by the size (2–2.5 3–3.5 mm) and shape (depressedovate) of the bract of staminate cymules, its most distinctive
characteristic. These remarkable bracts are markedly wider
than other species of the genus, which are lanceolate to linear,
0.5–1 mm). Other characterissometimes triangular (0.5–6
tics that distinguish the species are the shape and texture of
the basilateral glands of the bracts of staminate cymules,
which are sessile, ear-shaped, and creased. In other species of
this genus these glands are stipitate, rarely sessile, lanceolate,
and smooth. Additionally, in A. roselii the cataphylls that cover
the buds and persist at the base of the inflorescence are much
less developed than in the other species. An identification key
is herein presented aiming to facilitate the recognition of the
species of Actinostemon occurring in Brazil.
Actinostemon roselii is morphologically similar to A. concolor,
which is corroborated by the disposition and shape of foliar
glands, capsule globoid, pedicel accrescent, and persistent
calyx and stigma in the fruits. Although they are sympatric
species, they can be differentiated by the other aforementioned characteristics and also by the following traits: foliar
buds globoid (vs. ellipsoid in A. concolor), covered by cucullate
cataphylls (vs. cataphylls of varied shapes, oval to spatulate),
bract of staminate cymule depressed-ovate, slightly cucullate
(vs. lanceolate bract), glands of bracts of staminate cymules
ear-shaped, creased, and sessile (vs. globoid, smooth, and
stipitate), staminate flowers with (2) 3–4 (5) stamens (vs. 2–
10 stamens), and presence of sepals in staminate and pistillate
flowers (vs. achlamydeous flowers).
Distribution and Habitat—Actinostemon roselii is restricted
to the southern region of Brazil, in the states of Paraná (800–
1,000 m elevation) and Santa Catarina (50–590 m elevation)
(Fig. 2). It occurs in the domains of the Atlantic Forest, in
areas of ombrophilous forest and ecotonal areas between
ombrophilous dense forest and mixed ombrophilous forest,
frequently next to water bodies (riparian forests), in open
places, with the constant presence of Gymnanthes klotzschiana.
In Parque Estadual do Guartelá (Paraná), this species occurs in
areas of isolated woods (locally called capão) near the top of
the slope, in deeper soils, characteristic of montane mixed
ombrophilous forest. These areas of isolated woods (capão) are
generally connected to or involving riparian forests (IAP 2002).
Phenology—The species flowers from September to October
and bears fruits from October to February.
Etymology—The species epithet honors Dr. Roseli Barros,
a Brazilian botanist at Universidade Federal do Piauı́, who
introduced the first author to the science of plant taxonomy.
Conservation—Lack of detailed information about the
populations (e.g. size, fragmentation, etc.), ecology, area of
occupancy, and extension of occurrence places, makes A. roselii
is classified in the category of data deficient (DD) species
according to IUCN (2014).
Vernacular Name—One specimen cites the common name
as branquilho (voucher: Falkenberg & Albuquerque 4868).
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sepals and style column persistent; pedicel 2.7–3.8 cm long,
accrescent; columella 4–8 mm long, claviform. Seeds 3–4.2
2–2.2 mm, ellipsoid, smooth surface, sometimes slightly
granular, carunculate, dark brown, sometimes with blackish
spots. Figure 1.
[Volume 40
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Key to ACTINOSTEMON SPECIES OCCURRING IN BRAZIL
1.
Leaves grouped in the apex of branches conferring them a pseudoverticillate aspect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Cataphylls in the base of inflorescence villous. Panicle-cymose inflorescences. Ovary densely villous . . . . . . . . . . . . . . . . . . . . . . . . A. amazonicus
2. Cataphylls in the base of inflorescence glabrous to glabrescent. Racemose-cymose inflorescences. Ovary hirsute,
pubescent to glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2015]
OLIVEIRA ET AL.: A NEW SOUTH BRAZILIAN SPECIES OF ACTINOSTEMON
525
Fig. 2. Map showing the geographical distribution of Actinostemon roselii.
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3. Cataphylls in the base of inflorescence with veins little evident to absent. Leaves obovate, rarely elliptic.
Axis of inflorescences glabrous to glabrescent. Ovary glabrous to glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. appendiculatus
3. Cataphylls in the base of inflorescence with evident veins. Leaves elliptic to narrowly elliptic.
Axis of inflorescences pubescent to villous. Ovary hirsute, pubescent to glabrescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. verticillatus
Leaves regularly distributed along the branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Glabrous plants. Leaves of branches isophyllous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Ovary and fruits with echinate surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. echinatus
5. Ovary and fruits with smooth surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Foliar buds ellipsoid. Bract of staminate cymule 0.7–5.1 0.5–1.7 mm. Staminate cymule 3–8 flowers.
Staminate and pistillate flowers achlamydeous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. concolor
6. Foliar buds globoid. Bract of staminate cymule 2–2.5 3–3.5 mm. Staminate cymule 3–4(5) flowers.
Staminate flowers with 1–3 sepals, rarely absent; pistillate flowers with 3 sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. roselii
4. Indumented plants. Leaves of branches anisophyllous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Presence of leaves in the axis of the inflorescence. Stamens 6–12 in central flowers and 3–10 in lateral flowers . . . . . . . . . . . . . . A. klotzschii
7. Absence of leaves in the axis of the inflorescence. Stamens 5–7 in central flowers and 2–4 in lateral flowers . . . . . . . . . . . . . A. schomburgkii
Acknowledgments. The authors thank Conselho Nacional de
Desenvolvimento Cientı́fico e Tecnológico (CNPq 1413000/2011-0) and
the Post-graduate Program in Botany (PPGB) of the Universidade
Federal Rural de Pernambuco (UFRPE) for the grants; Coordenação
de Aperfeiçoamento de Pessoal de Nı́vel Superior (CAPES/PDSE
9208/12-7, CAPES/PNADB 23038000033/2010-16, and CAPES/PNPD)
for supporting the visits to international and national herbaria, and for
providing the postdoctoral grant, respectively; project REFLORA named
“Sistemática, filogenia e acervo virtual de coleções tipo de Euphorbiaceae,
com ênfase nas tribos Hippomaneae, Hureae e Crotoneae” (CNPq 563571/
2010-1) for the financial support; the curators of the herbaria where the
collections were analyzed, especially Gill Challen (K) and Dr. Hans-Joachim
Esser (M); Antônio Lins and Leidiana Lima for the map; and Frank Silva
for the illustration.
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